through time falls off as an exponential decay function; in other words, the percentage (but not the absolute number) of species going extinct in each period of time stays the same (Van Valen, 1973).1 These regularities, such as they are, have been interrupted during the past 250 million years by major episodes of extinction that have been recently estimated to occur regularly at intervals of 26 million years (Raup and Sepkoski, 1984).

Because of the relative richness of fossils in shallow marine deposits, the longevity of fish and invertebrate species living there can often be determined with a modest degree of confidence. During Paleozoic and Mesozoic times, the average persistence of most fell between 1 and 10 million years: that is, 6 million for echinoderms, 1.9 million for graptolites, 1.2 to 2 million for ammonites, and so on (Raup, 1981, 1984).

These estimates are extremely interesting and useful but, as paleontologists have generally been careful to point out, they also suffer from some important limitations. First, terrestrial organisms are far less well known, few estimates have been attempted, and thus different survivorship patterns might have occurred (although Cenozoic flowering plants, at least, appear to fall within the 1- to 10-million-year range). More importantly, a great many organisms on islands and other restricted habitats, such as lakes, streams, and mountain crests, are so rare or local that they could appear and vanish within a short time without leaving any fossils. An equally great difficulty is the existence of sibling species—populations that are reproductively isolated but so similar to closely related species as to be difficult or impossible to distinguish through conventional anatomical traits. Such entities could rarely be diagnosed in fossil form. Together, all these considerations suggest that estimates of the longevity of natural species should be extended only with great caution to groups for which there is a poor fossil record.

RAIN FORESTS AS CENTERS OF DIVERSITY

In recent years, evolutionary biologists and conservationists have focused increasing attention on tropical rain forests, for two principal reasons. First, although these habitats cover only 7% of the Earth’s land surface, they contain more than half the species in the entire world biota. Second, the forests are being destroyed so rapidly that they will mostly disappear within the next century, taking with them hundreds of thousands of species into extinction. Other species-rich biomes are in danger, most notably the tropical coral reefs, geologically ancient lakes, and coastal wetlands. Each deserves special attention on its own, but for the moment the rain forests serve as the ideal paradigm of the larger global crisis.

Tropical rain forests, or more precisely closed tropical forests, are defined as habitats with a relatively tight canopy of mostly broad-leaved evergreen trees

1

Van Valen’s original formulation, whose difficulties and implications are revealed by more recent research, has been discussed by Raup (1975) and by Lewin (1985). These studies deal with the clade, or set of populations descending through time after having split off as a distinct species from other such populations. They do not refer to the chronospecies, which is just a set of generations of the same species that is subjectively different from sets of generations.

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